Apples

The domesticated apple traces back to Malus sieversii, a wild apple species still found growing in the Tian Shan mountains of Kazakhstan, where wild apple forests produce fruit in a startling range of sizes, colors, and flavors.

Apples spread west along the Silk Road, picking up genetic contributions from other wild species along the way, eventually reaching Europe and, with colonization, North America. Nearly every apple variety grown today is a graft, not a seedling — apples don't reliably reproduce true from seed, so every "Honeycrisp" or "McIntosh" tree in the world is a genetic clone, propagated by grafting cuttings onto rootstock rather than growing from seed.

A striking number of major American apple varieties were quite literally accidents. McIntosh began in 1811 when John McIntosh, clearing land in Ontario, found wild seedlings and transplanted one that survived. Golden Delicious was found on Anderson Mullins's farm in West Virginia, sold to Stark Brothers Nursery for $5,000 in 1914. Granny Smith began as a chance seedling Maria Ann Smith discovered near a creek on her Australian property in 1868.

Honeycrisp has one of the more remarkable modern stories: the original tree, bred at the University of Minnesota in 1960, was badly winter-damaged in 1976–77 and written off as a failure, surviving only by accident through a few forgotten propagated clones. Breeder David Bedford rediscovered one in 1982 while reviewing trees marked for termination. DNA testing later revealed its true parentage (Keepsake and an unreleased breeding line called MN1627) was completely different from the Macoun-and-Honeygold story long repeated by nurseries.

Thomas Jefferson grew Esopus Spitzenburg and Newtown Pippin at Monticello, and Benjamin Franklin, while living in London in 1759, wrote home asking his wife to send him Newtown Pippins.

The University of Minnesota's apple breeding program, run for decades by Jim Luby and David Bedford, is responsible for Honeycrisp, Zestar! (released 1999, a cross of State Fair and breeding line MN 1691), and SweeTango (Honeycrisp × Zestar!, marketed exclusively through a controlled grower cooperative). DNA analysis found Duchess of Oldenburg, a hardy 19th-century Russian variety imported in 1835, present in the ancestry of 27 of the 28 cultivars the program has ever released — including both Wealthy and Honeycrisp.

The Purdue-Rutgers-Illinois (PRI) cooperative program, tracing back to early-20th-century crossing studies by C.S. Crandall at the University of Illinois and formalized in the 1940s, has released a series of scab-resistant apples bred using disease-resistance genes from a wild crabapple: Goldrush (1994), Enterprise (1993, one of the parents of the modern Cosmic Crisp), Liberty, and Pixie Crunch.

Cornell University's Geneva, New York station — the oldest apple breeding program in the US — released Jonagold (1968, Golden Delicious × Jonathan), Empire (1966, Red Delicious × McIntosh), Macoun, Cortland, and Freedom, a disease-resistant 1958 cross whose breeder, Robert Lamb, was honored decades later when Cornell named its newest 2020 release, Cordera, after him.

Because apples don't grow true from seed, every commercially grown apple tree is propagated by grafting a cutting from an existing tree onto rootstock — meaning every McIntosh tree on Earth is, genetically, the same individual plant as the original 1811 tree John McIntosh found.